dynamics and soils of oak/saw palmetto scrub after ftre ...nasa technical memorandum 103817...
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NASA Technical Memorandum 103817
Dynamicso-"-----_Vegetation and Soils ofOak/Saw Palmetto Scrub After Ftre:Observations From Permanent Transects
January 1991
(NASA-TM-IO3817) DYNAMICS OF VEGETATION ANO
SOILS OF OAK/SAW PALMETTO SC_U_ A=TER FIRE:
O_SERVATIONS FROM PERMANENT TRANSECTS
(NASA) 1_2 p CSCL 13_
G_/45
N91-21_35
N/ ANational Aeronautics and
Space Administration
https://ntrs.nasa.gov/search.jsp?R=19910012322 2020-03-09T23:21:38+00:00Z
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NASA Technical Memorandum 103817
Dynamics of Vegetation and Soils ofOak/Saw Palmetto Scrub After Fire:Observations From Permanent Transects
Paul A. Schmalzer, Ph.D.G. Ross Hinkle, Ph.D.
The Bionetics CorporationKennedy Space Center, Florida 32899
January 1991
National Aeronautics and
Space Administration
John F. Kennedy Space Center
NA.SA
Table of Contents
Section Page
Table of Contents ............................................................................................................... i
Abstract ............................................................................................................................... ii
List of Tables ..................................................................................................................... iv
Ust of Figures ................................................................................................................... vi
Acknowledgments ............................................................................................................ x
Introduction ........................................................................................................................ 1
Methods .............................................................................................................................. 2
Results ................................................................................................................................ 5
Discussion ....................................................................................................................... 29
Conclusions .................................................................................................................... 33
Uterature Cited ............................................................................................................... 35
Appendix I. Species composition of scrub stands preburn through 36 monthspostbum ................................................................................................... 41
Appendix II. Chemical characteristics of scrub soils before and after theDecember 1986 fire ............................................................................... 52
Appendix II1. Ordination, community structure, and soil chemistry figures .......... 65
Abstract
Ten permanent 15 m transects previously established in two oak/saw
palmetto scrub stands burned in December 1986, while two transects remained
unburned. We sampled vegetation in the > 0.5 m and < 0.5 m layers on these
transects at 6, 12, 18, 24, and 36 months postburn and determined structural
features of the vegetation (height, % bare ground, total cover). We analyzed
vegetation data from each sampling by height layer using detrended
correspondence analysis ordination. Vegetation data for the > 0.5 m layer for
the entire time sequence were combined and analyzed using detrended
correspondence analysis ordination. Soils were sampled at 6, 12, 18, and 24
months postburn and analyzed for pH, conductivity, organic matter,
exchangeable cations (Ca, Mg, K, Na), NO3-N, NH4-N, AI, available metals (Cu,
Fe, Mn, Zn), and PO4-P.
Shrub species recovered at different rates postfire with saw palmetto
reestablishing cover >0.5 m within one year, but the scrub oaks had not
returned to prebum cover >0.5 m in 3 years after fire. These differences in
growth rates resulted in dominance shifts after fire with saw palmetto increasing
relative to the scrub oaks. Such shifts were not uniform across the scrub
gradient. Mixed oak/saw palmetto-dominated transects changed the most and
recovered more slowly than either oak or saw palmetto-dominated transects.
Distances between preburn and successive postburn locations in ordination
space appear to be a useful index of this recovery. In mature scrub, ordination
of the > 0.5 m layer reflected the environmental gradient from wet to dry.
Ordinations of the >0.5 m layer from 6 through 18 months after fire reflected the
rate of recovery as well as environmental differences, while ordinations of the <
0.5 m layer reflected the main environmental gradient. Most of the forbs and
grasses in scrub also resprouted after fire; several of these were more common
in the wetter transects. Overall changes in species richness were minor,
although changes occurred in species richness by height layers due to different
growth rates. Mean total cover > 0.5 m increased linearly after fire but did not
reach a maximum in 3 years. Mean total cover < 0.5 m was 50% at 6 months
postfire and remained that at 36 months postfire. Mean height was about 76 cm
at 3 years postfire.
Soils of well drained and poorly drained sites differed markedly. Soil
responses to fire appeared minor. Soil pH increased at 6 and 12 months
postfire; calcium increased at 6 months postburn. Nitrate-nitrogen increased at
12 months postburn. Low values of conductivity, PO4-P, Mg, K, Na, and Fe at 12
months postburn may be related to heavy rainfall the preceeding month.
Seasonal variability in some soil parameters appeared to occur.
I11
List of Tables
Table 1.
Table 2.
Table 3.
Table 4.
Table 5.
Table 6.
Table 7.
Table 8.
Table 9.
Table I-1.
Table I-2.
Table !-3.
Table I-4.
Table II-1.
Page
Composition (percent cover) of scrub stands that burned inDecember 1986 pre- and postbum (> 0.5 m) ................................. 6
Composition (percent cover) of scrub stands that burned inDecember 1986 pre- and postbum (< 0.5 m) ................................. 8
Composition (percent cover) of transects in scrub Stand 1 thatburned in December 1986 pre- and postburn (> 0.5 m) ............. 11
Composition (percent cover) of transects in scrub Stand 1 thatburned in December 1986 pre- and postburn (< 0.5 m) ............. 13
Composition (percent cover) of transects in scrub Stand 2 thatburned in December 1986 pre- and postburn (> 0.5 m) ............. 15
Composition (percent cover) of transects in scrub Stand 2 thatburned in December 1986 pre- and postburn (< 0.5 m) ............. 17
Composition of unburned scrub transects in Stand 1 after theDecember 1986 fire ........................................................................... 19
Differences between preburn and postburn axis 1 ordinationscores from detrended correspondence analysisordination ............................................................................................ 24
Euclidean distances between preburn and postburn locations(axis 1, axis 2) of transects in ordination space from detrendedcorrespondence analysis ordination .............................................. 26
Composition (percent cover) of scrub stands that burned inDecember 1986 prebum .................................................................. 42
Composition (percent cover) of scrub stands that burned inDecember 1986 at 6 and 12 months postbum ............................. 44
Composition (percent cover) of scrub stands that burned inDecember 1986 at 18 and 24 months postburn .......................... 47
Composition (percent cover) of scrub stands that burned inDecember 1986 at 36 months postburn ........................................ 50
Chemical characteristics of well drained scrub soils where thevegetation burned in December 1986 before and after thefire ......................................................................................................... 53
iv
Table 11-2.
Table 11-3.
Chemical characteristics of poorly drained scrub soils where thevegetation burned in December 1986 before and after thefire ......................................................................................................... 57
Chemical characteristics of well drained soils where thevegetation remained unburned before and after the December1986 fire ............................................................................................... 61
V
List of Figures
Figure 1.
Figure 2.
Figure 3.
Figure 4.
Figure 5.
Figure 6.
Figure 7.
Figure 8.
Figure 9.
Figure 10.
Figure 11.
Page
Detrended correspondence analysis ordination of the > 0.5 mlayer of scrub transects sampled in January 1985 that burned inDecember 1986 .................................................................................... 67
Detrended correspondence analysis ordination of the < 0.5 mlayer of scrub transects sampled in January 1985 that burned inDecember 1986 .................................................................................... 69
Detrended correspondence analysis ordination of the > 0.5 mlayer of scrub transects that burned in December 1986 sixmonths postbum ................................................................................... 71
Detrended correspondence analysis ordination of the < 0.5 mlayer of scrub transects that burned in December 1986 sixmonths postbum ................................................................................... 73
Detrended correspondence analysis ordination of the > 0.5 mlayer of scrub transects that burned in December 1986 12 monthspostbum .................................................................................................. 75
Detrended correspondence analysis ordination of the < 0.5 mlayer of scrub transects that burned in December 1986 12 monthspostbum .................................................................................................. 77
Detrended correspondence analysis ordination of the > 0.5 mlayer of scrub transects that burned in December 1986 18 monthspostbum .................................................................................................. 79
Detrended correspondence analysis ordination of the < 0.5 mlayer of scrub transects that burned in December 1986 18 monthspostbum .................................................................................................. 81
Detrended correspondence analysis ordination of the > 0.5 mlayer of scrub transects that burned in December 1986 24 monthspostbum .................................................................................................. 83
Detrended correspondence analysis ordination of the < 0.5 mlayer of scrub transects that burned in December 1986 24 monthspostbum .................................................................................................. 85
Detrended correspondence analysis ordination of the > 0.5 mlayer of scrub transects that burned in December 1986 36 monthspostbum .................................................................................................. 87
vi
Figure 12.
Figure 13.
Figure 14.
Figure 15.
Figure 16.
Figure 17.
Figure 18.
Figure 19.
Figure 20.
Figure 21.
Figure 22.
Figure 23.
Figure 24.
Detrended correspondence analys_sordination of the < 0.5 mlayer of scrub transects that burned in December i986 36 monthspostbum..................................................................................................89
Detrended correspondence analysts sample ordination of the >0.5 m layer of all burned transects of scrub vegetation frompreburn through 36 monthspostburn...............................................91
Detrended correspondence analysasspecies ordination of the >0.5 m layer of all burned transects of scrub vegetation fromprebum through 36 monthspostburn...............................................93
Detrended correspondence analys_s sample ordination of the >0.5 m layer showing changes of selected transects from preburnthrough 36 months postbum ............................................................... 95
Detrended correspondence analysis sample ordination of the >0.5 m layer showing changes of selected transects from preburnthrough 36 months postburn ............................................................... 97
Detrended correspondence analys_s sample ordination of the >0.5 m layer showing changes of selected transects from preburnthrough 36 months postbum ............................................................... 99
Mean total cover prebum and through 36 months postburn of the> 0.5 m layer of scrub transects that burned in December1986 ...................................................................................................... 101
Mean total cover preburn and through 36 months postburn of the< 0.5 m layer of scrub transects that burned in December1986 ...................................................................................................... 103
Percent bare ground preburn and through 36 months postburn ofscrub transects that burned in December 1986 ............................ 105
Mean height preburn and through 36 months postburn of scrubtransects that burned in December 1986 ....................................... 107
Mean maximum height preburn and through 36 months postburnof scrub transects that burned in December 1986 ....................... 109
Species richness (mean number of species per transect) preburnand through 36 months postburn of the > 0.5 m layer of scrubtransects that burned in December 1986 ....................................... 111
Species richness (mean number of species per transect) preburnand through 36 months postburn of the < 0.5 m layer of scrubtransects that burned in December 1986 ....................................... 113
vii
Figure 25.
Figure 26.
Figure 27.
Figure 28.
Figure 29.
Figure 30.
Figure 31.
Figure 32.
Figure 33.
Figure 34.
Figure 35.
Figure 36.
Species richness (mean number of species per transect) preburnand through 36 months postburn of both strata of scrub transectsthat burned in December 1986 ........................................................ 115
Hydrogen ion concentration of well drained scrub soils preburnand through 24 months postburn of scrub transects that burned inDecember 1986 .................................................................................. 117
pH of well drained scrub soils preburn and through 24 monthspostburn of scrub transects that burned in December 1986 ...... 119
Conductivity of well drained scrub soils preburn and through 24months postbum of scrub transects that burned in December1986 ...................................................................................................... 121
Organic matter of well drained scrub soils preburn and through 24months postbum of scrub transects that burned in December1986 ...................................................................................................... 123
Available phosphorus of well drained scrub soils preburn andthrough 24 months postburn of scrub transects that burned inDecember 1986 .................................................................................. 125
Exchangeable calcium of well drained scrub soils preburn andthrough 24 months postburn of scrub transects that burned inDecember 1986 .................................................................................. 127
Exchangeable magnesium of well drained scrub soils preburnand through 24 months postburn of scrub transects that burned inDecember 1986 ................................................................................. 129
Exchangeable potassium of well drained scrub soils preburn andthrough 24 months postburn of scrub transects that burned inDecember 1986 .................................................................................. 131
Exchangeable sodium of well drained scrub soils preburn andthrough 24 months postburn of scrub transects that burned inDecember 1986 .................................................................................. 133
Exchangeable nitrate-nitrogen of well drained scrub soils preburnand through 24 months postburn of scrub transects that burned inDecember 1986 .................................................................................. 135
Exchangeable ammonium-nitrogen of well drained scrub soilspreburn and through 24 months postburn of scrub transects thatburned in December 1986 ................................................................ 137
viii
Figure 37.
Figure 38.
Figure 39.
Figure 40.
Figure 41.
Figure 42.
Exchangeable aluminum of well drained scrub soils preburn andthrough 24 months postburn of scrub transects that burned inDecember 1986 .................................................................................. 139
Available copper of well drained scrub soils preburn and through24 months postburn of scrub transects that burned in December1986 ...................................................................................................... 141
Available iron of well drained scrub soils prebum and through 24months postbum of scrub transects that burned in December1986 ..................................................................................................... 143
Available manganese of well drained scrub soils preburn andthrough 24 months postburn of scrub transects that burned inDecember 1986 .............................................................. . ................... 145
Available zinc of well drained scrub soils preburn and through 24months postburn of scrub transects that burned in December1986 ...................................................................................................... 147
Precipitation from December 1986 to December 1988 during thecourse of the scrub soil study ........................................................... 149
ix
Acknowledgments
This study was conducted under NASA contracts NAS10-10285 and
NAS10-11624. We thank William M. Knott, II1, Chief, Biological Research and
Ufe Support Office, Albert M. Koller, Jr., formerly, Chief, Programs and Planning
Office, and Burt Summerfield, Pollution Control Officer, Biomedical Operations
and Research Office, for their guidance. We thank Lee Maull and Joseph
Mailander for field assistance, Teresa Englert and Steve Black for chemical
analysis, and Opal Tilley and Julie Harvey for manuscript preparation.
X
Introduction
Scrub vegetation dominated by shrub oaks (Quercus chapmanii, Q.
geminata, Q. myrtifolia), ericaceous shrubs (e.g., Lyonia fruticosa, L. lucida) and
saw palmetto (Serenoa repens) is an important upland vegetation type on
Kennedy Space Center (KSC) (Provancha et ai. 1986). it is a major habitat for
the threatened Florida scrub jay (Aphelocoma coerulescens coerulescens)
(Breininger 1981, 1989) and is used by the gopher tortoise (Gopherus
polyphemus) and other species of concern (Breininger et al. 1988).
Oak/saw palmetto scrub vegetation is fire adapted and recovers from fire
primarily by sprouting (Abrahamson 1984 a,b, Schmalzer and Hinkle 1987).
Previously, we examined four stands of scrub vegetation of differing ages on
similar sites in the Happy Creek area of KSC, established permanent transects,
and resampled those transects in the absence of fire (Schmalzer and Hinkle
1987). The youngest stand sampled had burned two years before the first
sampling.
In December 1986, a fire burned through two of the stands previously
sampled. Resampling after this fire was conducted to determine early postfire
responses not covered in the previous study. In addition, there are advantages
to examining changes on permanent sample plots or transects rather than
inferring temporal changes from sites of differing ages but similar in other
environmental conditions (Mueller-Dombois and Ellenberg 1974, Austin 1977).
Examples of permanent plot studies in Florida vegetation include Veno (1976),
Abrahamson (1984a,b), Givens et al. (1984), Myers (1985), and Menges (1990).
The study site is an inland area of Merritt Island. Merritt Island has a
warm, humid climate. Annual precipitation averages 131 cm, but year-to-year
variability is high. Precipitation varies seasonally with a wet season occurring
from May to October and the rest of the year being relatively dry (Mailander
1990). Thunderstorms are frequent in the summer months with lightning strikes
being common (Eastern Space and Missile Center 1989). Moisture deficits
typically occur between mid-March and mid-May and between mid-November
and mid-December (Mailander 1990). Mean daily maximum temperatures are
22.3oc for January and 33.3oc for July; mean daily mininum temperatures are
9.6oc for January and 21.9oc for August (Titusville records, Mailander 1990).
The scrub stands in this study occur primarily on Pomello sand (Arenic
Hapiohumod), a moderately well drained soil, but the wetter end of the scrub
gradient is on the poorly drained Immokalee (Arenic Haplaquod) or Myakka
sand (Aeric Haplaquod) (Huckle et ai. 1974). These soils are low in available
nutrients with much of the nutrient standing crops in living and dead vegetation
and litter rather than the mineral soil (Schmalzer and Hinkle 1987).
Methods
Vegetation Sampling and Analysis
Permanent vegetation transects were established and sampled in four
scrub stands in January 1983; in January 1985, they were resampled
(Schmalzer and Hinkle 1987). The December 1986 fire burned through scrub
Stands 1 and 2; Stands 3 and 4 did not burn. In Stand 1 four of six transects
(#2, 3, 4, 5) burned, and in Stand 2 all six transects (#7-12) burned. We
2
sampled vegetation of the burned transects at 6, 12, 18, 24, and 36 months
postfire using line-intercept techniques and sampling the 0-0.5 m and > 0.5 m
height layers (Muelller-Dombois and Ellenberg 1974). Unburned transects
(Stands 1, #1 and 6) were sampled 6, 24, and 36 months postfire. Sampling
methods used were the same as before the fire.
Vegetation data from each sampling period were analyzed by height
layer using detrended correspondence analysis ordination (Hill and Gauch
1980, Gauch 1982) in the PCORD package (McCune 1987). Vegetation data
from all sampling periods were combined and detrended correspondence
analysis ordination was used with this larger data set to examine the patterns of
compositional change. Ordination techniques have been used previously to
study vegetation dynamics (Austin 1977, Whittaker and Woodwell 1978,
Swaine and Greig-Smith 1980, Menges 1990), and recovery after fire (Hobbs
and Gimingham 1984, Westman and O'Leary 1985, Malanson and Trabaud
1987). The data from repeated sampling of the same scrub transects appeared
particularly suited to this approach.
Soil Sampling and Analysis
All transects in Stand 1 were on soils mapped as Pomello sand
(moderately well drained); in Stand 2, 4 transects (#9-12) were on Pomello
sand and 2 transects (#7, 8) were on Myakka sand (poorly drained) (Huckle et
al. 1974).
We sampled soils from the 0 to 15 cm and 15 to 30 cm depths near each
burned transect immediately postfire and 6, 12, 18, and 24 months postfire.
Unburned transects were sampled 6, 12, and 24 months postfire. Each sample
was a composite of at least 5 soil cores. Soil samples were homogenized and
3
subsampled; subsamples were oven-dried at 50o C for 24 hours. Oven-dried
samples were analyzed for nitrate-nitrogen and ammonium-nitrogen. Other
analyses were conducted on air-dried samples.
Analyses of all parameters except organic matter were made in the
NASNKSC Environmental Chemistry Laboratory. Soil pH was determined on a
1:1 soil to water slurry (McLean 1982) using an Orion pH meter. Conductivity
was measured on a 1:5 soil to water solution using a conductivity meter
(Rhoades 1982). Exchangeable cations, Ca, Mg, Na, and K, were extracted in
neutral 1N ammonium acetate (Knudsen et al. 1982, Lanyon and Heald 1982)
and analyzed by atomic absorption spectrophotometer (Perkin-Elmer
Corporation 1982). Available metals, copper (Cu), iron (Fe), manganese (Mn),
and zinc (Zn), were extracted in diethylenetrlaminepentaacetic acid (DTPA)
(Olson and Ellis 1982, Gambrell and Patrick 1982, Baker and Amacher 1982)
and analyzed by atomic absorption spectrophotometer. Exchangeable
aluminum was extracted in 1N potassium chloride (Barnhisel and Bertsch 1982)
and analyzed by atomic absorption spectrophotometry (Perkin-Elmer
Corporation 1982). Exchangeable nitrate-nitrogen (NO3-N) and ammonium-
nitrogen (NH4-N) were extracted in 2N potassium chloride (Keeney and Nelson
1982) and then analyzed on a Technicon Autoanalyzer (Technicon Industrial
Systems 1973, 1983a). Available phosphorus was determined by extraction in
deionized water (Olsen and Sommers 1982) followed by analysis on a
Technicon Autoanalyzer (Technicon Industrial Systems 1983c). Organic matter
was determined by the combustion method (Nelson and Sommers 1982).
Organic matter determinations were made by Post, Buckley, Schuh, and
Jernigan, Inc., Orlando, Florida.
4
Results
Vegetation Composition
The response of all burned transects (Table 1, Table 2, Appendix I,
Tables I-1 to I-4) indicated regrowth of the shrub species of scrub at differing
rates post/ire. In the > 0.5 m layer (Table 1), cover of saw palmetto returned to
preburn values in one year after fire, wax myrtle (Myfica cefifera) required 2
years to reestablish preburn cover, Ilex glabra, Befafia racemosa, and Lyonia
lucida required 3 years. The scrub oaks had not returned to preburn cover by 3
years after the fire. Myrtle oak (Quercus myrtifolia) had only about half its
preburn cover by 3 years postfire.
In the < 0.5 m layer (Table 2), there were increases after fire in the cover
of shrubs that before burning were in the • 0.5 m layer. These included Lyonia
lucida and L. fruticosa, Ilex glabra, Myrica cerifera and the scrub oaks. By 3
years after fire, cover of many of these shrubs < 0.5 m was beginning to decline
as they grew into the > 0.5 m layer. Some subshrubs (Vaccinium myrsinites,
Gaylussacia dumosa) that often do not reach 0.5 m height also increased after
the fire.
Response of herbaceous species varied. Perennial grasses such as
Aristida stficta and Andropogon spp. resprouted and reestablished cover. There
appeared to be modest increases in cover of Carphephorus spp., Panicum spp.,
and Eupatorium rotundifo/ium. Pteridium aqui/inum and Ga/actia e//iottii
appeared only in the 6 months and 18 months postburn samples. This was a
seasonal effect, since both species are deciduous. The fire occurred in
5
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summer conditions.
The two stands that burned were not identical preburn. Stand 1 was
older (11 vs. 7 years since fire) and on a drier site than Stand 2 (see Schmalzer
and Hinkle 1987). Therefore, we also considered their postfire response
separately. Stand 1 (Table 3, Table 4, see also Appendix I) showed the same
general trends discussed above except that some shrubs such as Iiex glabra
did not occur on the drier sites, and there were fewer herbaceous species.
Conversely, Stand 2 (Table 5, Table 6, see also Appendix I) followed the same
general trends but had additional shrub and herbaceous species.
The two unburned transects in Stand 1 (Table 7) showed only minor
changes in composition and percent cover in the time since the December 1986
fire.
Ordination Analysis
Preburn ordination of the > 0.5 m layer (Appendix III, Figure 1) gave a
pattern where Transects 7 and 8 defined the wet end of the gradient dominated
by saw palmetto with ilex glabra and Persea borbonia, while the oaks
dominated the xeric end. The < 0.5 m layer gave a similar ordination pattern
(Appendix III, Figure 2), reflecting the depth to water table gradient. Transect 7
was not represented in this ordination, since preburn it had no vegetation < 0.5
m.
At 6 months postbum, ordination of the > 0.5 m layer (Appendix I11,Figure
3) had Transect 5 at one end and the other transects clustered near the other
end of the first axis. Immediately after fire, regrowth of saw palmetto exceeded
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that of the oaks and dominated the > 0.5 m layer. Transect 5 lacked saw
palmetto, and its regrowth was dominated by the oaks, as indicated in the
species ordination. In contrast, the < 0.5 m layer ordination (Appendix III, Figure
4) reflected the environmental gradient previously shown in the > 0.5 m layer.
At 12 months postburn, Transect 5 still defined one end of the
ordination of the > 0.5 m layer (Appendix III, Figure 5). More species had grown
into this layer, including some of those that characterize wetter sites. The < 0.5
m layer ordination (Appendix III, Figure 6) reflected the depth to water table
gradient with Transects 7 and 8 defining the wet end.
By 18 months postfire (Appendix !11,Figure 7), Transect 5 still defined the
oak-dominated end of the first axis of the > 0.5 m layer ordination, Transects 7
and 8 emerged as saw palmetto-dominated transects on the other end, and a
group of transects with mixed dominance was intermediate. Ordination of the <
0.5 m layer (Appendix III, Figure 8) reflected the differences between Transects
7 and 8 and the others. In species composition, the wet end of the gradient was
partly determined by herbaceous species (e.g., Panicum spp., Eupatorium
rotundifolium) and small shrubs (e.g., Fthus copailina), since other shrubs had
grown into the > 0.5 m layer.
At 24 months postburn, the overall gradient was well defined in the > 0.5
m layer (Appendix !11,Figure 9), but less so in the < 0.5 m layer (Appendix III,
Figure 10). The ordination patterns at 36 months postburn were similar to those
24 months postburn (Appendix III, Figures 11, 12). Comparing the preburn
ordination of the > 0.5 m layer (Appendix III, Figure 1) to 36 months postburn
(Appendix III, Figure 11) indicated a general but incomplete recovery of the
community pattern by this time. Oak cover was still less than preburn in the > 0.5
21
m layer; this was reflected in the intermediate location of many transects that
preburn had greater oak dominance as indicated by their closer location to
Transect 5 in the preburn ordination.
Further details of the recovery process are given by the detrended
correspondence analysis ordination of the transects at all sampling times. The
overall pattern (Appendix i11,Figure 13) had the oak-dominated transects (e.g.,
#5) to the right side of the ordination and saw palmetto-dominated transects
(#7,#8) to the left, as is also indicated by the species ordination (Appendix III,
Figure 14).
Trajectories of individual transects over time are seen more easily by
plotting fewer of the transects on one graph (Appendix III, Figures 15, 16, 17).
Three patterns of responses occurred. Transect 5 with high oak dominance and
little saw palmetto cover remained on the right side of the ordination (Appendix
II!, Figure 15). At 6 months after fire, it was located higher on the second axis,
probably due to the cover of Galactia and Smilax in that sample. Over time, the
position of the transect in the ordination space tracked back toward its original
location but was not identical to prebum by 36 months postburn.
Saw palmetto-dominated transects (#7, Appendix !11,Figure 16; #8,
Figure 15) remained on the left side of the ordination. At 6 months postburn,
both of these transects were located higher on the second axis than preburn,
probably reflecting the cover of Eupatorium and Pteridium postfire. Transect 7
tracked back toward its preburn location relatively smoothly. Transect 8 showed
a variation in that the 18 months postburn sample was closer to the 6 months
than the 12 months sample. This was a seasonal response, since the herbs
22
Eupatorium and Pteridium were represented in the summer (6, 18 months)
samples but not the winter ones.
Most of the transects (# 2, 3, 4, 9, 10, 11, 12) were originally of mixed
oak/saw palmetto dominance or oak dominated but with saw palmetto
abundant. These all moved from the oak end of the ordination to the saw
palmetto end after fire and then gradually tracked back toward their original
location (Appendix III, Figure 15, #10, 11, 12; Figure 16, #2, 4; Figure 17, #3, 9).
Transect 2 showed a slight variation in this pattern in that the sample 12 months
postburn had greater saw palmetto dominance than at 6 months after the fire;
this seems to be due to slower initial growth on this transect.
We calculated two indices intended to reflect recovery based on the
ordination results. The first was simply the difference between the first axis
ordination score of the sample taken before the fire and the first axis score for
each postfire sampling of that transect. The first axis was used since this is the
most important in an ordination. These differences declined with increasing time
since fire (Table 8). The group of transects of mixed dominance showed the
greatest change 6 months postfire and the largest remaining differences 36
months postfire compared to the saw palmetto-dominated transects or the the
single oak-dominated one (Table 8).
The second index was the euclidean distance between the preburn
location of a transect and its location at each postbum sampling based on the
first two ordination axes. This index takes into account information from the
second ordination axis. See Malanson and Trabaud (1987) for a similar
approach. These values declined with increasing time after the fire and
23
Table 8. Differences Between Preburn and Postburn Axis 1 Ordination Scores FromDetrended Correspondence Analysis Ordination.
6 Months 12 Months 18 Months 24 Months 36 MonthsTransect Postburn Postburn Postburn Postburn Postburn
2 62 86 39 38 213 113 94 59 55 204 106 85 72 67 14
5 36 38 9 11 07 20 22 13 21 7
8 10 7 13 1 (-2) 1
9 84 74 46 12 (-12) 110 96 95 70 45 1411 140 133 97 90 4112 49 3O 22 8 2
--2
Xai i 71.6 66.4 44.0 34.8 13.3
2
SDal I 43.0 40.0 30.1 29.3 12.2
-3
Xsp 15.0 14.5 13.0 11.0 4.5
3
SDsp 7.1 10.6 0 14.1 3.5
-4
Xo/sp 92.9 85.3 57.9 45.0 17.7
4
S Do/sp 31.0 30.6 24.7 29.2 12.0
1 Calculations based on absolute values of negative differences.2 All transects (N--10).
3 Saw palmetto transects (#7, 8) (N=2).
4 Oak/saw palmetto transects (#2, 3, 4, 9, 10, 11, 12) (N=7).
24
• alsosuggested that the mixed dominance transects changed to a greater
degree than the saw palmetto or oak-dominated ones (Table 9).
Vegetation Structure
Mean total cover > 0.5 m increased almost linearly from 6 to 36 months
postfire (Appendix III, Figure 18). Mean total cover < 0.5 m (Appendix III, Figure
19) reached a value of about 50% at 6 months postfire and fluctuated within that
range through 36 months postfire. Percent bare ground (Appendix III, Figure 20)
was high immediately after fire but declined rapidly, approaching zero at 36
months postfire. Mean height (Appendix II!, Figure 21) increased rapidly through
one year after fire and then at a slower rate. Mean maximum height (Appendix
III, Figure 22) increased rapidly to 6 months after fire and then changed slowly.
Species dchness > 0.5 m (mean number of species per transect) (Appendix III,
Figure 23) increased to 18 months postfire. Species dchness < 0.5 m (Appendix
I!1, Figure 24) and species richness of all strata (Appendix i11,Figure 25) reached
high values at 6 months postfire and changed little from then through 36 months
postfire.
Soil Chemistry
Previous work (Schmalzer and Hinkle 1987) and initial analysis of the
postburn soil data indicated that the more poorly drained scrub soils had higher
levels of organic matter and nutrients related to organic matter. In order to
reduce variation within the soil data set, well drained soils (8 burned transects)
were considered separately from the poorly drained ones (2 burned transects).
Some data from 2 unburned transects were also collected. The sample sizes of
the poorly drained and unburned soils were not sufficient for definite
conclusions. Soils of these transects were sampled previously in January 1983.
25
Table 9. Euclidean Distances Between Preburn and Postburn Locations (Axis 1, Axis2) of Transects In Ordination Space from Detrended CorrespondenceAnalysis Ordination.
6 Months 12 Months 18 Months 24 Months 36 MonthsTransect Postburn Postburn Postburn Postburn Postburn
2 70.7 102.6 46.3 41.2 22.13 130.4 113.7 63.3 56.5 20.64 114.0 101.8 83.4 75.6 •15.25 81.4 42.9 20.1 11.0 29.07 35.2 33.3 25.6 21.8 9.28 81.6 43.6 83.0 15.0 3.69 104.4 88.8 56.6 12.2 15.01 0 104.8 101.6 70.1 45.5 50.01 1 155.2 148.9 114.1 96.6 54.61 2 49.0 31.0 22.0 8.2 9.2
-- 1
Xal I 92.7 80.8 58.5 38.4 22.9
1
SDai I 36.5 40.4 30.8 30.4 17.2
-2
Xsp 58.4 38.5 54.3 18.4 6.4
2
SDsp 32.8 7.3 40.6 4.8 4.0
-3
Xo/sp 104.1 98.3 65.1 48.0 26.7
3
S Do/sp 35.5 35.3 29.0 31.9 18.1
1 All transects (N=10).2 Saw palmetto transects (#7, 8) (N=2).
3 Oak/saw palmetto transects (#2, 3, 4, 9, 10, 11, 12) (N=7).
26
These data are given in Appendix II, Tables I1-1 to 11-3.Graphs are presented
only for the well drained, burned soils.
Hydrogen ion concentration (Appendix Iii, Figure 26) decreased at 6 and
12 months after the fire and then returned to preburn values by 18 months
postburn. Soil pH (Appendix III, Figure 27) increased with the hydrogen ion
decrease and then returned to preburn values. Poorly drained soils (Table 11-2)
showed decreased hydrogen ion concentration (increased pH) at 12 and 24 but
not 18 months postburn; unburned soils (Table 11-3) showed a decrease in
hydrogen ion concentration at 12 months after the fire.
Conductivity declined at 12 months postburn (Appendix I11,Figure 28).
This decline occurred in the burned, poorly drained scrub (Table 11-2) but also in
the unburned scrub (Table 11-3).
Organic matter (Appendix !11,Figure 29) did not show a clear pattern of
change after the fire. At 12 months postburn, values were low in the surface
layer, but 18 and 24 month postburn values were not. Poorly drained soils
(Table 11-2) and unburned soils (Table 11-3) did not show this trend. As
previously shown, organic matter levels were much higher in poorly drained
than well drained soils.
Phosphorus (Appendix III, Figure 30) did not show a definite fire
response. Values at 12 months postbum were lower than those preceeding and
following. The same trend occurred in the poorly drained soils (Table 11-2).
Calcium appeared slightly elevated at 6 months postfire (Appendix III,
Figure 31) but by 12 months postburn returned more in range with preburn
values. Magnesium (Appendix II1, Figure 32) did not show a definite fire
27
response. As with several other parameters, it decreased at 12 months postfire
but increased later. Potassium (Appendix III, Figure 33) decreased at 12 months
postburn, but values at 18 and 24 months postburn were higher than preburn.
The poorly drained soils (Table 11-2)also displayed this pattern. Sodium
(Appendix iil, Figure 34) showed a slight decrease at 12 months postburn
followed by an increase at 18 and 24 months. Poorly drained soils (Table 11-2)
were similar; unburned soils (Table 11-3)also had higher values 24 months
postfire than before the fire.
Nitrate-nitrogen (Appendix I!!, Figure 35) was low immediately and 6
months postburn, increased at 12 months postburn, and remained high at 18
and 24 months postbum. The poorly drained soils (Table !1-2)also displayed
this pattern. Unburned soils (Table 11-3)increased at 24 but not 12 months
postf]re. Ammonium-nitrogen (Appendix III, Figure 36) declined between
immediately postburn and 12 months postburn, then increased sharply at 18
months, declining again at 24 months postburn. Poorly drained soils (Table 11-2)
followed the same pattern. Variability was evident even in the unburned soils
(Table 11-3).
Aluminum (Appendix i11,Figure 37) increased at 18 months postburn in
well drained soils and in poorly drained ones (Table 11-2).Unburned soils also
increased in aluminum at 24 months postfire (Table 11-3).Copper (Appendix I11,
Figure 38) increased at 18 months postburn in well drained soils and in poorly
drained ones (Table 11-2),but unburned soils changed little (Table 11-3).Iron
(Appendix 111,Figure 39), manganese (Appendix I11,Figure 40), and zinc
(Appendix I!1,Figure 41) varied some through time but did not show clear
changes from fire.
28
Discussion
Vegetation Composition and Structure
Regrowth of oak/saw palmetto scrub immediately after fire was primarily
by the resprouting of shrubs present before the fire, as indicated in our previous
work (Schmalzer and Hinkle 1987) and in studies of similar vegetation
(Abrahamson 1984a,b). Rates of regrowth differed between species. This is
seen most clearly immediately after fire; saw palmetto reestablished preburn
cover > 0.5 m in one year after fire, while scrub oaks did not reestablish preburn
cover > 0.5 m in 3 years. Abrahamson (1984a,b) observed similar species
differences in postfire regrowth.
Mean total cover > 0.5 m three years postfire was similar to cover
recorded in our earlier study (Schmalzer and Hinkle 1987). At two years
postfire, cover was greater than in the two year old stand previously sampled;
that stand had greater oak dominance and exhibited slower reestablishment of
cover in the > 0.5 m layer. Mean total cover < 0.5 m was also in the range
reported previously (Schmalzer and Hinkle 1987). The decline in cover < 0.5 m
that occurred in stands 4 years old (Schmalzer and Hinkle 1987) did not occur
by 3 years postfire here.
Differential growth rates of the major shrub species resulted in shifts in
dominance after fire. Ordination analyses indicated that these shifts were not
uniform but were most pronounced in transects of mixed dominance. While
earlier studies (Schmalzer and Hinkle 1987) indicated dominance changes,
these changes were most pronounced in the first two years after fire, as
reflected here.
29
Herbaceous species such as Pteridium and Eupatofium increased after
fire, particulary in the saw palmetto-dominated transects, but this was of much
less magnitude than the shrub response. Much of the herbaceous response
was vegetative regrowth rather than seedling establishment. This was certainly
the case with grasses (Andropogon, Afistida) and perennial forbs such as
Ptefidium and Galactia. Some species (e.g., Eupatorium) were encountered
primarily in summer sampling; since sampling before the fire (Schmalzer and
Hinkle 1987) was conducted in the winter, seedling establishment cannot be
ruled out. At most, however, this is a minor component of the vegetation
recovery in oak/saw palmetto scrub.
The subshrubs Vaccinium myrsinites and Gaylussacia dumosa
increased immediately after fire. Abrahamson (1984a,b) observed this in Lake
Wales Ridge vegetation but noted that these shrubs can persist for lengthy
pedods without fire.
A consequence of the dominance of sprouting species in oak/saw
palmetto scrub vegetation and the rapid recovery of plant cover after fire is that
there is little change in overall species richness. There are changes in species
richness by strata due to differing rates of height growth. This confirms earlier
results (Schmalzer and Hinkle 1987). Abrahamson (1984a) noted that only a
small set of plant species were well-adapted to the set of envirionmental
conditions that characterize scrub; these conditions include winter drought,
acid, nutrient-poor, sandy soils, and repeated burning. In rosemary (Ceratiola
ericoides) scrub, openings persist longer after fire due to the slower recovery by
a seeder species, and there was a greater postfire response by herbaceous
species (Johnson and Abrahamson 1990). Fire is thought to maintain species
30
diversity in many shrublands (Gill and Groves 1981, Kruger 1983, Christensen
1985).
Bare ground did not persist long after fire in the oak/saw palmetto scrub.
This is consistent with our previous study (Schmalzer and Hinkle 1987) and
with observations that there is little bare ground in undisturbed oak/saw
palmetto scrub not recently burned (Breininger et ai. 1988, Breininger and
Schmalzer 1990). Oak/saw palmetto scrub differs from much sand pine scrub
(Mulvania 1931, Webber 1935) where openings are common and persist.
Rates of height growth were similar to those reported before (Schmalzer
and Hinkle 1987). Mean height was 76 cm, well below one meter, at 3 years
after fire.
Soil Chemistry
In general, values of most soil parameters were similar to previous
results (Schmalzer and Hinkle 1987). For all major nutrients, concentrations
were greater in the 0-15 cm layer than at 15-30 cm. Poorly drained scrub soils
were higher in organic matter and most nutrients than the well drained soils.
There are several complications to interpreting the responses to fire of
these soils. The preburn samples were taken 3 years before the fire, not
immediately preburn. There was not a matched sampling of soils from unburned
scrub; therefore, seasonal variation of soil parameters may influence the results.
Madsen (1980) conducted quarterly sampling of KSC soils including those from
scrub but concluded that the data were not sufficient to establish seasonal
trends.
31
The decrease in hydrogen ion concentration (pH increase) at 6 and 12
months postfire is probably due to fire. Soil pH increases of differing
magnitudes are commonly observed after fire and are related to the release of
cations in ash (Raison 1979, Wells et al. 1979). The only cation to show an
increase coinciding with pH is calcium which is the most abundant cation in
scrub biomass and litter (Schmalzer and Hinkle 1987). Abrahamson (1984a)
observed an increase in calcium after fire that lasted less than one year.
Conductivity, phosphorus, magnesium, potassium, sodium, and iron all
decreased at 12 months postburn. By 18 and 24 months postburn all returned in
line with preceeding values or, in the case of sodium and potassium, exceeded
previous levels. Precipitation records (Appendix II!, Figure 42) indicated that the
rainfall of November 1987 (9.44 in, 24.0 cm), the month preceeding the 12
months postburn sampling, was the greatest monthly precipitation during the
course of the study. It is possible that this resulted in leaching of mobile
elements from the upper layers of the soil.
Nitrate-nitrogen and ammonium-nitrogen exhibited delayed responses to
fire. Nitrate-nitrogen began increasing at 12 months extending to 18 and 24
months postburn. Similar delayed increases in nitrate-nitrogen occurred after
fire in coastal strand vegetation on Canaveral National Seashore (Hinkle et al.
1989) and vadous other systems including chaparral (Christensen and Muller
1975, DeBano et al. 1977), South African fynbos (Stock and Lewis 1986), and
pocosins (Wilbur and Christensen 1983). Delayed increases in nitrate-nitrogen
have been related to initial declines in nitrifying bacteria (Dunn and DeBano
1977) followed by their subsequent recovery and mineralization of nitrogen
32
made available by fire (DeBano et al. 1979, Raison 1979). Total Kjeldahl
nitrogen data from these sites are not yet available.
Reasons for the increase of aluminum and copper at 18 and 24 months
postburn are unclear. Increases in pH decrease the availability of aluminum
and copper (Brady 1974, Marschner 1986), but the magnitude of the pH change
here appears insufficient to have much effect. Additionally, pH returned to its
preburn values at 18 months postburn, while aluminum and copper
considerably exceeded preburn values then.
Conclusions
1. The general pattern of recovery of oak/saw palmetto scrub after fire is
that of sprouting by the dominant shrub species with little overall change in
species composition or species richness. Sampling of permanent transects
repeatedly after fire reveals details of this process which were not as clear from
previous work. Recovery patterns differ over the scrub gradient. Dominance of
mixed oak/saw palmetto transects changes much more immediately after fire
than either oak-dominated or saw palmetto-dominated transects, and recovery
is not complete by 3 years after fire.
2. Many of the structural changes after fire are persistent. Percent bare
ground is reduced quickly by resprouting shrubs, but cover in the > 0.5 m layer
and shrub height recover more slowly. Yet unresolved are potential changes
from repeated, frequent fires that might reduce root carbohydrate reserves (e.g.,
Hough 1968, Harrington 1985, 1989).
33
3. Scrub soils do not appear greatly changed by fire. There is a modest
increase in pH that last for about one year after fire and an increase in calcium.
There is a delayed increase in nitrate-nitrogen as seen in several other studies.
Low values of several parameters at 12 months after the fire may be related to
unusually high rainfall the preceeding month. Future work in effects of fire on
scrub soils needs to examine the broader question of fire effects on nutrient
cycling in scrub including losses of nutrients directly from fire (e.g., Raison et al.
1985a,b), possible importance of gaseous emissions of nitrogen oxides
(Anderson et al. 1988, Levine et al. 1988, 1990), and the potential for leaching
losses. Soil studies may require balanced sampling of unburned sites to
account for seasonal variability which appears greater than anticipated.
34
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38
Nelson, D.W. and L.E. Sommers. 1982. Total carbon, organic carbon, andorganic matter. In: A.L. Page, R.H. Miller, and D.R. Keeney (eds.).Methods of soil analysis, part 2. Chemical and microbial properties.Agronomy 9:539-579. American Society of Agronomy, Inc., Madison,Wisconsin.
Olsen, S.R. and L.E. Sommers. 1982. Phosphorous. In: A.L. Page, R.H. Miller,and D.R. Keeney (eds.). Methods of soil analysis, part 2. Chemical andmicrobial properties. Agronomy 9:403-430. American Society ofAgronomy, Inc., Madison, Wisconsin.
Olson, R.V. and R. Ellis, Jr. 1982. Iron. In:A.L. Page, R.H. Miller, and D.R.Keeney (eds.). Methods of soil analysis, part 2. Chemical and microbialproperties. Agronomy 9:301-312. American Society of Agronomy, Inc.,Madison, Wisconsin.
Perkin-Elmer Corporation. 1982. Analytical methods for atomic absorptionspectrophotometry. Perkin-Elmer Corporation. Norwalk, Connecticut.
Provancha, M.J., P.A. Schmaizer, and C.R. Hinkle. 1986. Vegetation types. JohnF. Kennedy Space Center, Biomedical Operations and Research Office(Maps in Master Planning format, 1:9600 scale, digitization by ERDAS,Inc.).
Raison, P.J. 1979. Modification of the soil environment by vegetation fires, withparticular reference to nitrogen transformations: a review. Plant and Soil51:73-108.
Raison, R.J., P.K. Khanna, and P.V. Woods. 1985a. Mechanisms of elementtransfer to the atmosphere during vegetation fires. Canadian Journal ofForest Research 15:132-140.
Raison, R.J., P.K. Khanna, and P.V. Woods. 1985b. Transfer of elements to theatmosphere during low-intensity prescribed fires in three Australiansubalpine eucalypt forests. Canadian Journal of Forest Research 15:657-664.
Rhoades, J.D. 1982. Soluble salts. In: A.L. Page, R.H. Miller, and D.R. Keeney(eds.). Methods of soil analysis,part 2. Chemical and microbialproperties. Agronomy 9:167-179. American Society of Agronomy, Inc.,Madison, Wisconsin.
Schmalzer, P.A. and C.R. Hinkle. 1987. Effects of fire on composition, biomass,and nutrients in oak scrub vegetation on John F. Kennedy Space Center,Florida. NASA Technical Memorandum 100305. John F. Kennedy SpaceCenter, Florida. 146pp.
39
Stock, W.D. and O.A.M. Lewis. 1986. Soil nitrogen and the role of fire as amineralizing agent in a South African coastal fynbos ecosystem. Journalof Ecology 74:317-328.
Swaine, M.D. and P. Greig-Smith. 1980. An application of principal componentsanalysis to vegetation change in permanent plots. Journal of Ecology68:33-41.
Technicon Industrial Systems. 1973. Nitrate and nitrite in water and wastewater.Industrial method no. 100-70W. Technicon Industrial Systems. Tarrytown,New York.
Technicon Industrial Systems. 1983a. Ammonia in water and wastewater. In:Multi-test cartridge method no. 696-82W: 1D-4D. Technicon IndustrialSystems, Tarrytown, New York.
Technicon Industrial Systems. 1983b. Ortho-phosphorous. In: Multi-testcartridge no. 696-82W: 1B-4B. Technicon Industrial Systems, Tarrytown,New York.
Veno, P.A. 1976. Successional relationships of five Florida plant communities.Ecology 57:498-508.
Webber, H.J. 1935. The Florida scrub, a fire-fighting association. AmericanJournal of Botany 22:344-361.
Wells, C.G., R.E. Campbell, LF. DeBano, C.E. Lewis, R.L Fredriksen, E.C.Franklin, R.C. Froelich, and P.H. Dunn. 1979. Effects of fire on soil. USDAForest Service General Technical Report WO-7. 34pp.
Westman, W.E. and J.F. O'Leary. 1986. Measures of resilience: the response ofcoastal sage scrub to fire. Vegetatio 65:179-189.
Whittaker, R.H. and G.M. Woodwell. 1978. Retrogression and coenoclinedistance. Pp. 51-70. In: R. H. Whittaker (ed.). Ordination of plantcommunities. Dr. W. Junk. The Hague, The Netherlands.
Wilbur, R.B. and N.L Christensen. 1983. Effects of fire on nutrient availability ina North Carolina coastal plain pocosin. American Midland Naturalist110:54-61.
4O
Appendix I
Species Composition of Scrub Stands Preburn Through 36 Months Postburn
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51
Appendix II
Chemical Characteristics of Scrub Soils Before and After the December 1986
Fire
52
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64
Appendix I!i
Ordination, Community Structure, and Soil Chemistry Figures
65
Figure 1. Detrended correspondence analysis ordination of the > 0.5 m layer of
scrub transects sampled in January 1985 that burned in December 1986.
Sample ordination shows individual transects. Species shown are ARISTR
(Aristida stricta), BEFRAC (Befaria racemosa), ILEGLA ( llex glabra), LY©FRU
(Lyonia fruticosa), LYOLUC (Lyonia lucida), MYRCER (Mynca cerifera),
PERBOR (Persea borbonia), QUECHA ( Quercus chapmanil), QUEGEM
( Quercus geminata), QUEMYR ( Quercus myrtifolia), and SERREP ( Serenoa
repens).
66
100
DCA ORDINATIONPREBURN
GREATER THAN 0.5 m
(N
Or}I
X<
P-5
75
5O
25
0 L0 25
rl
P-IOOO
P-4P-ll
0 P-3
r'l P-2
P-9
l I0 I
50 75
(-I P-12
100
AXIS I
I
125
P-8 []
P-7 []
t I
150 1 75 200
250
SPECIES
("4
X<
200
150
100
5O
0
-50
-100
-150 -
-200-50
El LYOFRU
0 QUECEM
[] QUECHA
[] C)UZMYR
13 ARISTR
0 BEFRAC
I_ LYOLUC
0 MYRCER
0 SERREP
I 1 I
0 50 100 150
AXIS
.17
2OO 250
ILEGLA 0
PERBOR
[]I
300 35O
Figure 2. Detrended correspondence analysis ordination of the < 0.5 m layer of
scrub transects sampled in January 1985 that burned in December 1986.
Sample ordination shows individual transects. Species are as in Figure 1 with
the addition of HYPRED (Hypericum reductium), SMIAUR (Smilax aurfculata),
and VACMYR (Vaccinium myrsinites).
68
150
DCA ORDINATIONPREBURN
LESS THAN 0.5 m
¢N
(/3m
X
125
100
75
50
25
0
I=-11
P-2
O
0
0 P-5
I50
[] P-3
[] P-4
[] P-IO
I-1 P-9
I I
P-8
P-t2
[
[]
1O0 150 200 250
AXIS 1
500
SPECIES
¢'4
X
4.00
300
2OO
100
0
-100 -
-200 -
-100
[] SMIAUR
C] L.YOFRU
[] QUECHA I-1 QUEWYR
[] UYRCER
[]
[] VACMYR
LYOLUC [][] QUEGEM
0 BEFRAC
SERREP0
I
HYPRED
[] ARISTR
0I I t I
2OO
AXIS 1
,39
100 300
ILEGLA
4-00
[]
Figure 3. Detrended correspondence analysis ordination of the > 0.5 m layer of
scrub transects that burned in December 1986 six months postburn. Species
are as previously defined with the addition of EUPROT (Eupatorium
rotundifolium), GALELL (Galactia elliottil), and PTEAQU (Ptefidium aquilinum).
7o
100
DCA ORDINATION6 MONTHS POSTBURNGREATER THAN 0.5 m
¢M
(/1u
x
75
5O
25
0
P-10
P-4P-11
0
-_ P-7
P-9
P-8
0
I
25
O P-2
O P-12
I
5O1 I I
75
AXIS I
100 125
P-5O
150
SPECIES
¢'4
O'3
X
3OO
200
100
0
--100
-200
-300
_/3 LYOFRU
- _ EUPROT
[] M_C_
O $ERREP
[] LYOLUCO C_LELL
[] QUEMYR
I I I I I
SMIAUR
[] QUEGEM
O
-50 0 50 100
AXIS 171
150 200 250
Figure 4. Detrended correspondence analysis ordination of the < 0.5 m layer of
scrub transects that burned in December 1986 six months postburn. Species
are as previously defined with the addition of BAREGR (Bare ground), CARSPP
(Carphephorus spp.), GAYDUM (Gaylussacia dumosa), LICMIC (Licania
michauxil), PANSPP (Panicum spp.), RHUCOP (Rhu$ copallina), and VACSTA
( Vaccinium stamineum).
72
150
6DCA ORDINATIONMONTHS POSTBURNLESS THAN 0.5 rn
¢M
(/1!
X
125
100
75
5O
25
0
[] P-12
O P-10
P-11r'l
P-3 P-5
I P-2
0
P-4.0
I-IP-9
I
5O
0 P-8
I I I
I"IP-7
1O0 150 200 250
AXIS 1
300
SPECIES
O4
u
X<
250
200
150
100
50
0
-50
-100
-150
-200
VACSTA0 SERREP
O QUECE'M
BAR[GR0
0 AItlSTR
0 LYOFRU
[] CARSPP
OUEMYR VACMYR0
[] Q UCMIC
O LYOLUC
r'l PANSPPQUECHA
r_ I_ SMIAUR
OIGAYOUM I I I I
O0 -50 0 50
O GALELL
I
2OO100
AXIS?3
150
0 B_m.A,¢
_YRCF.R0
PTEAQU
I
250
0HYPRED
ILEGLA0
0
EUPROTO
RHUCOP
300
Figure 5. Detrended correspondence analysis ordination of the > 0.5 m layer of
scrub transects that burned in December 1986 12 months postburn. Species
are as previously defined.
"74
75
DCA ORDINATION12 MONTHS POSTBURNGREATER THAN 0.5 m
C,4
(/3
X<
5O
25
0
P-8
P-11m
EJ P-lO
0 P-7
P-9O
0
P-,I.O
P-2
[] OP-3
I
25
r'l P-12
I I
50 75
AXIS 1
I
100
0 P-5
I
125
350
SPECIES
("4
x<
,3OO
250
200
150
100
5O
0
-50
-100
-150
[] ILEGLA
[] MYRCER
0 PERBOR
I
O0 -50
13LYOFRU
[] SERREP
LYOLUC
I 0
0
0 BEFRAC
I I
50 100
AXIS 175
[] QUECEM
QUEMYRl-I
O QUECHA
I I I
150 200 250
ARtSTR[3
3OO
Figure 6. Detrended correspondence analysis ordination of the < 0.5 m layer of
scrub transects that burned in December 1986 12 months postburn. Species
are as previously defined with the addition of ANDSPP (Andropogon spp.) and
DROSPP (Drosera spp.).
76
125
DCA ORDINATION12 MONTHS POSTBURN
LESS THAN 0.5 m
¢,,I
m
x.<
100
75
5O
25
0
P-5[]
_P--11
0 P-4
[] p-3
D P-2
I 1
0 25 50
[] P--IO
[] P-12
P-OI -o I I I
75 1O0 125 150
AXIS 1
0 P-7
I
175
oP-8
2OO
X.<
250
200
150
100
5O
0
-50
-100
-150
-200
-250
-300
__SMIAUR
[] UCMIC
- DROSPP
D [] LYOFRU
O QUECHA
- O QUEMYR
D GAYDUM
1 I
O0 -50 0
SPECIES
u
QUEGEM
[] BAREGR
rq VACMYR
CARSPP [] MYRCER
BEFRAC
0 LYOLUC
O SERREP EUPROT
PANSPP []
[]
[] ILEGLA
O ANDSPP
HYPRED
[] ARISTR
I I I
50 100 150
AXIS9"7
2OO
1
I 1 I
250 300 350 4.00
Figure 7. Detrended correspondence analysis ordination of the > 0.5 miayer of
scrub transects that burned in December 1986 18 months postburn. Species
are as previously defined.
78
tO0
DCA ORDINATION18 MONTHS POSTBURNGREATER THAN 0.5 m
(/1I
X
75
5O
25
0
P-8
P-4[]
P-7
0 P-12
[] P-11
O P-9
[] P-3
P-10 P-2I [] I 0 1
0 25 50 75
I
100
AXIS
I I
125 150
0 P-5
1
175 200
300
SPECIES
-e,,l
(/3x<
250
200
150
100
50
0
-50
-100
-150
-200I
[] ILEGLA
PERBOR[]
RHUCOP
i I
50 - IO0
I
-50
LYOFRU
0 0 SERREP[] GALELL
El LYOLUC
I-IQUECHA
0 ARIS'TR
MYRCER
0 0 BEFRAC
1 I I 1
5O
AXIS
79
100 150 200
QUEGEM[]
QUEMYR[]
250
Figure 8. Detrended correspondence analysis ordination of the < 0.5 m layer of
scrub transects that burned in December 1986 18 months postburn. Species
are as previously defined with the addition of SEYPEC (Seymeria pectinata)
and UNKHER (Unknown herb).
8o
200
DCA ORDINATION18 MONTHS POSTBURN
LESS THAN 0.5 m
IN
m
x
150
100
5O
0
[] P-12
0 P-lO
[] P-4_-11 [] P-3
I0 P-2
I
0 50I I
1O0 150
AXIS
P-8[]
I
200
I
250
P-7 []
300
350
300 O SEYPEC
SPECIES
IN
m
x
250
200
150
100
5O
0
-50
-100
-150
I QUEGEM
D °_- VACSTA
L D SERREP
[] LYOFRU
n ARISlt_[]BAREGR
i
- [] VACMYR
O QUEMYR
O QUECHA
I I I
-50 0 50
0 CARSPP
[] LYOLUC
UCMIC[] SMIAUR
0 MYRCER
[] ANDSPP
[] GALELL
PANSPP[]
HYPRED ILEGLA
0 O0
PTEAQU
GAYDUM
I [] I
EUPROT[]
RHUCOP
UNKHER
100 150 200
AXIS 181
I I I I
250 300 350 4-00
Figure 9. Detrended correspondence analysis ordination of the > 0.5 m layer of
scrub transects that burned in December 1986 24 months postburn. Species
are as previously defined.
82
125
DCA ORDINATION24 MONTHS POSTBURNGREATER THAN 0.5 m
Or)X<1:
IO0
75
5O
25
[] P-7
P-8
00
0 P-4
I-/ P-9
P-3[] C] P-2
C] P-12
P-I 1 [] P-10
I [] I I I I I
25 50 75 100 125 150
AXIS 1
n P-5
175
5OO
SPECIES
(/1x
4.O0
300
2OO
100
0
-100
-200
[] ANDSPP
[] ARISTR
I-1 PE'RBOR
ILEGLA
[3 LYOLUC
rl SERREP
[] QUECHA[]QUEMYR
[] QUEGEM
I
-100
MYRCER 0
LYOFRUL []
0
[] BEFRAC
I I I I
1O0
AXIS
83
200
1
300
SMIAUR []
400
Figure 10. Detrended correspondence analysis ordination of the < 0.5 m layer
of scrub transects that burned in December 1986 24 months postburn. Species
are as previously defined.
i
i
! 84
125
DCA ORDINATION24 MONTHS POSTBURN
LESS THAN 0.5 m
¢,,I
m
x<
100
75
5O
25
0
I"1 P-4
] P-11
rl P-lO
0 P-5
0 P-9
P-3[]
[] P-2
I I
0 5O 1O0
0 P-12
0 P-8
I I I
1,50 2O0 25O
AXIS 1
I
300
P-7 0
35O
300
SPECIES
x.<
2OO
IO0
0
-100
-200
-300
-400
-500
0LYOFRU
0 GAYDUM
-0 QUEMYR
QUECHA
[] [] BAREGR
0 ARIST'R
[] VACMYR
0 OROSPPl l
0 QUEGEM
0 LYOLUCMYRCERO
HYPRE.D0
[] SERREP
ANOSPP
I I I L I I
PANSPP
o%
ILEGLA
-5O 0 5O 100 1,50
AXIS
85
2OO
1
250 300 350 4-OO
Figure 11. Detrended correspondence analysis ordination of the > 0.5 m layer
of scrub transects that burned in December 1986 36 months postburn. Species
are as previously defined.
86 C.-_
150
DCA ORDINATION,36 MONTHS POSTBURNGREATER THAN 0.5 rn
x
125
100
75
5O
25
0
P-5
0
I
25
I
5O
[] P-IO
I
75
O P-3
P-9
O O P-2
[] P-4
[] P-11
P-12
o I100
AXIS 1
1 I I125 150 175
O P-8
P-7 []
20O
SPECIES
(/3
X.<
3OO
250
200
150
100
5O
0
-50
-100
-150 1-200
AI_ISTR 0
0 QUEGEM
I0
O BEFRAC
0 MYRCER0LYOFRU
O QUECHA
0 QUEMYR
LYOLUC
00 SERREP
ILEGLA0
[]
PERBOR
I I 1100
AXIS 187
2OO 300 4.00
Figure 12. Detrended correspondence analysis ordination of the < 0.5 m layer
of scrub transects that burned in December 1986 36 months postburn. Species
are as previously defined.
1O0
DCA ORDINATION,56 MONTHS POSTBURN
LESS THAN 0.5 m
¢N
X
75
50
P-11L
i
25
0
0
P-2
(3P-¢
I
50
P-5[]
Q P-3O
I100
O P-8
I-1 P-12
P-10 [] I"I P-g
I I150 200
AXIS 1
I
250
O P-7
I300 35O
SPECIES
w
x
300
250
200
150
100
50
0 -
-50 -
-100 -
-150 -
-200 -
-250-50
QUEMYRO
O MYRC_
[] r-I VACMYRDROSPP
LYO_UO QUECHA O
[] GAYDUM
[] CARSPP
0 BAREGR
1 1 L__
0 5O 100 150
I"ILYOLUC
n ARISTR
/
2OO
AXIS89
BEFRAC
J-4a-----_250 300
1
n HYPRED
iI F'G_[]
O ANDSPP
O PANSPP
.1 J_
350 400 4-5O
Figure 13. Detrended correspondence analysis sample ordination of the > 0.5
m layer of all burned transects of scrub vegetation from preburn through 36
months postburn.
9O
r-_0_
0
00cN
o
o
o
o
o
o
o
0
SIXV
91
Figure 14. Detrended correspondence analysis species ordination of the > 0.5
m layer of all burned transects of scrub vegetation from preburn through 36
months postburn. Species are as previously defined.
92
g
WOO(-DObJo_>
m
<[ I
£DZn.r_-O
Fn
CD CL[]
OO
O
[]
[]
I
CDOW9
Or,- _.0. o _S
0 0W _ tl_ w
•1- 0[] _. m -
r. [][] _rn
I t ...... I
0 0 00 004 ,--
1 l ! I
O O O O OO O O O O_- ("4 W') _- LD
I I I I I
Z SIXY
OOuU
OO
OOw9
c_ or)
O
OO
I
93
Figure 15. Detrended correspondence analysis sample ordination of the > 0.5
m layer showing changes of selected transects from preburn through 36 months
postburn. Shown are transects 5, 8, 10, 11, and 12.
94
m
I--
r-_n
C_-- O 0_- -- -- CO u') _;
I i i i i _.O O En o. i1. I:L n o i_
oEO E
+_I14, O { O_.
"- O C_I
o_ o_ _'*
!; +_ 2° _ •
o _°E _
o _ _ _0
.L l 1 L J
0
E(LC)rO
C) C) O C) O O C) OLO ,_ ("q C) 00 (.O ,_- ('N
OC:)oq
O00
O(.C)
O
C)(-,q
O° 09i
X<1:
O00
C)(D
O
O(N
O
C)
glX_
95
Figure 16. Detrended correspondence analysis sample ordination of the > 0.5
m layer showing changes of selected transects from preburn through 36 months
postburn. Shown are transects 2, 4, and 7.
96
m
_rn
WU3O0Wn>
03
r_ Z_0
0
I I I13.0.0..
uu_o
0 0cD '_-
0E
0E
0 _"E e_0o0 O
0o E oE _ DE
"- <I
0
E []
_ o o _E E _
i I'- o •
<I mm__mm
I I I I -- in
0 0 0 0O,4 0 O0 CO
Z: SIXV
0E
cOI'0
[]
o&
I
0
1
0o4
(23(23o4
0- 00
0- '43
0
0- 0'4
!
"- X
000
0£{3
0
00,4
97
Figure 17. Detrended correspondence analysis sample ordination of the > 0.5
m layer showing changes of selected transects from preburn through 36 months
postburn. Shown are transects 3 and 9.
98
m
I--:D.,_mI--LIJ (.fl(-DO1113_>.
OO
rY z(Do
m
i--
i:_ZFyrl/0
rn
I In n
oE
I I I
0E
£N 0"- E
O O O O(.13 '_" o4 O
OE
Q,,
4>
O
O
o _
<><>
I I I I
O O O OcO _O _ eq
(:D(Do4
O- O0
O- r..O
O
O- O,I
_o_m
'- X.,<
O00
OqD
O
OO4
O
O
SIXV
9£.
Figure 18. Mean total cover preburn and through 36 months postburn of the >
0.5 m layer of scrub transects that burned in December 1986. Shown are
means and 95% confidence intervals. Data are the sum of the cover of
individual species per transect.
i00
_E
O0
_-0_W
Z_-
WW
0
0C_
I I
0 0•- 0
0
[]
I 0 I
m
I [] I
I I I ! I I I I
0 0 0 0 0 0 0 0O_ _ I_. _0 1.0 _" 1"9 (N
I'0
0
- 0
..0
n
!
0 0
W
oc0
( ) 3AO0
I01
Figure 19. Mean total cover preburn and through 36 months postburn of the <
0.5 m layer of scrub transects that burned in December 1986. Shown are
means and 95% confidence intervals. Data are the sum of the cover of
individual species per transect.
102
r_wEOur')
(_C)
(/').,_ u')i,IW
CDCD
I I I I
0 0 0 00") O0 I_ (0
I D Im
w
m
g
m
- C)
m
I--Q--4
I 1 I I I
0 0 0 0 0
CD
0
I.C)Cxl
E
i,i
1°n0
103
Figure 20. Percent bare ground preburn and through 36 months postburn of
scrub transects that burned in December 1986. Shown are means and 95%
confidence intervals.
104
0
0rY(.9
I,In/
133
I,IrjrY,i,I13_
00
I 1, 1 I I I I I I
0 0 0 0 0 0 0 0 0(_ O0 I_ _ I._ _'- I_ (N ,--
(_) aNn0a0 3av8
m
H3_
Ei
O
uO(N
c"O
-_ Ev
bJ
_o O
- uO
- O
_E
n
O
105
Figure 21. Mean height preburn and through 36 months postbum of scrub
transects that burned in December 1986. Shown are means and 95%
confidence intervals. Data are heights measured at four intervals (0, 5, 10, 15
m) along each transect.
106
cD
(.9m
W
<W
rn
r"FcO(.,r)
I I I
0 0 0 0
i..--..-.-C-........-_
_---....C).....--4
I--.-..!_-......-_
W !.- ! ! !
0 0 0 0 00") 00 I'-- (.0 i._
0 0 CD 0'_- r,') cN
(u o) ±HOI3H
_Ou
O
u3CN
O
EV
W
-° O<
- uD
- O
r_
0_
(D
107
Figure 22. Mean maximum height preburn and through 36 months postburn of
scrub transects that burned in December 1986. Shown are means and 95%
confidence intervals. Data are the maximum height of any shrub along each
transect.
108
(.9m
I,I
m_
<I:i,i
I E]
I I I
0 0 0 0eq 0 00
I
I
0cO
I I I I I I I
0 0 0 0 0 0 0C_ 0 O0 cO _ c_l
(_ ::;) J.HOI3H
O
uOr'O
Om
I'O
LDC,,I
_o_c:t-O
-_ Ev
i,i_o O
- O
1°.Dt,_r_
O
109
Figure 23. Species richness (mean number of species per transect) preburn
and through 36 months postburn of the > 0.5 m layer of scrub transects that
burned in December 1986. Shown are means and 95% confidence intervals.
ii0
E
L_Z o
rY-r
WrY
I0
I [] I
I [] I
I D I
I D I
I I I I I I I I I
SS3NHOIW $3103dS
0
uO_D
0i
i
V
W
- u'_
- CD
Q_
CD
iii
Figure 24. Species richness (mean number of species per transect) preburn
and through 36 months postburn of the < 0.5 m layer of scrub transects that
burned in December 1986. Shown are means and 95% confidence intervals.
112
-_Wuo
-ro(D
<
_)OOL_OOO-I__OO--I
D
I [] I
I [] I
I D I
I O I
I I I
I'Q ¢xl -- 0
I O I
I I I I I I I I I
O
u
_ O
cO
O
E
<
- u_
- O
..a
O_
SS3NHOIW "$3133dS
113
Figure 25. Species richness (mean number of species per transect) preburn
and through 36 months postburn of both strata of scrub transects that burned in
December 1986. Shown are means and 95% confidence intervals.
114
(.AW--Ii
W
t.A
I [] I
I 0 I
I i-! i
I [] I
I I ! I I I
0
uOI'D
0I,D
uO
E
W
- uO
- 0
1°$3103dSSS3NHOIW
115
Figure 26. Hydrogen ion concentration of well drained scrub soils preburn and
through 24 months postburn of scrub transects that burned in December 1986.
Shown are means and 95% confidence intervals for the 0-15 cm layer (A) and
the 15-30 cm layer (B).
116
20.000
18.000
16.000
14.000
)
D 12.000
X 10.000
8.000r
6.000
4.000
2.000
0.000
//
D
I
n
w
' //Prebum
Hydrogen IonA. 0-15 cm
0
I i
6 12
AGE (months)
!
18
i
24
14.000 //
B. 15-30 cm
12.000
10.000
- 8.000
=¢
r6.000
4.000
0.000 , //Prebum
I I I
0 6 12
AGE (months)117
i I
18
I
24
ORIGINAL PAGE IS
OF POOR QUALITY
Figure 27. pH of well drained scrub soils preburn and through 24 months
postburn of scrub transects that burned in December 1986. Shown are means
and 95% confidence intervals for the 0-15 cm layer (A) and the 15-30 cm layer
(B).
118
5.0 //,
pHA. 0-15 cm
-1-a.
4.5
4.0
3.5 , //Prebum
I I I I I
0 6 12 18 24
AGE (months)
5.0 //
B. 15-30 cm
"1"
4.5
4.0
3.5 , //Preburn
!
0I I
6 12
AGE (months)119
I I
18 24
Figure 28. Conductivity of well drained scrub soils preburn and through 24
months postburn of scrub transects that burned in December 1986. Shown are
means and 95% confidence intervals for the 0-15 cm layer (A) and the 15-30 cm
layer (B).
120
E(,3
U'}
oE-1
V
i,-m
!
I-,.0
Z0£.)
100
9O
8O
7O
60
5O
4O
3O
//
i //Preburn
CONDUCTIVITYA. 0-15 cm
I
0I
6 12
AGE (months)
I I
18 24.
E(J
oE
v
i-ra
m
I--(..):)
Z00
60
50
,¢0
30
20
10
,//
, //Preburn
I
0
B. 15-,50 cm
I 1
6 12
AGE (months)121
I 1
18 24
Figure 29. Organic matter of well drained scrub soils preburn and through 24
months postburn of scrub transects that burned in December 1986. Shown are
means and 95% confidence intervals for the 0-15 cm layer (A) and the 15-30 cm
layer (B).
122
I0.0 ,//
ORGANIC MATTERA. 0-15 cm
9.0
_,j 8.0
¢1:L_ 7.0I--I--
_E 6.0
__ 5.0-(.9r,,, 4.00
3.0
i2.0 i
rl
, //Prebum
I
0I I I
186 12
AGE (months)
I
2¢
3.0 //
B. 15-.30 cm
2.5
v
I:1:: 2.0LIJk--I--
_E 1.5
0
Z_C 1.0C.9
C)0.5
0.0 1
0
I 1 I
186 12
AGE (months)123
1
2+
Figure 30. Available phosphorus of well drained scrub soils preburn and
through 24 months postburn of scrub transects that burned in December 1986.
Shown are means and 95% confidence intervals for the 0-15 cm layer (A) and
the 15-30 cm layer (B).
124
6.0
5.0
_, 4..0
_t
3.0v
n 2.0
1.0
0.0
3.0
2.5
v
n 1.0
0.5
0.0
//
I IIPreburn
//
, //Preburn
I
0
I
0
PHOSPHORUSA. 0-15 cm
I I
6 12
AGE (months)
1 I
18 24.
B. 15-30 cm
I I I I
6 12
AGE (months)125
18 24.
Figure 31. Exchangeable calcium of well drained scrub soils preburn and
through 24 months postburn of scrub transects that burned in December 1986.
Shown are means and 95% confidence intervals for the 0-15 cm layer (A) and
the 15-30 cm layer (B).
126
350.0
300.0
-_ 250.0t_
.X
Ev
150.0
o
100.0
50.0
0.0
120.0
100.0
80.0
E 60.0
{3 40.00
20.0
0.0
//
m
- 0
o
, //Preburn
//
m
r
, //Prebum
I
0
CALCIUMA. 0-15 cm
1
0
I I
6 12
AGE (months)
I
18
B. 15-30 cm
I I I
186 12
AGE (months)12"7
I
2¢
I
2¢
Figure 32. Exchangeable magnesium of well drained scrub soils preburn and
through 24 months postbum of scrub transects that burned in December 1986.
Shown are means and 95% confidence intervals for the 0-15 cm layer (A) and
the 15-30 cm layer (B).
128
120.0
Ioo.o
O_"_ 80.0
20.0
//
o.o , //Preburn
30.0 //
25.0
O_ 20.0
O>
E 15.0
5.0
0.0 L
i"1
i //Preburn
MAGNESIUMA. 0-1,5 cm
I I i0 6 12
AGE (months)
I18
B. 15-30 cm
I
0
k
6
AGE
12
(months)I29
I
18l
24
Figure 33. Exchangeable potassium of well drained scrub soils preburn and
through 24 months postbum of scrub transects that burned in December 1986.
Shown are means and 95% confidence intervals for the 0-15 cm layer (A) and
the 15-30 cm layer (B).
130
90.0
80.0
70.0
"_ 60.0_h(
"_ 50.0
Ev 4.0.0
v 30.0
20.0
10.0
0.0
B
m
//
i //Prebum
I
0
POTASSIUMA. 0-15 cm
....... I I
6 12
AGE (months)
I
18
I
24.
50.0 //
B. 15-30 cm
t_E
v
v
4.0.0
30.0
20.0
10.0
0.0 , //Preburn
I
0I I
6 12
AGE (months)
I
18I
24.
131
Figure 34. Exchangeable sodium of well drained scrub soils preburn and
through 24 months postburn of scrub transects that burned in December 1986.
Shown are means and 95% confidence intervals for the 0-15 cm layer (A) and
the 15-30 cm layer (B).
132
60.0
50.0
40.0
E 30.0
0 20.0Z
10.0
0.0
50.0
40.0
_30.0
EV
20.0
(3Z
10.0
0.0
//
B
D
, //Prebum
//
R
B
I
0
SODIUMA. 0-15 cm
I I
6 12
AGE (months)
B. 15-30 cm
I
18
I
0
I I
6 12
AGE (months)
133
I
18
I
24
!
24
Figure 35. Exchangeable nitrate-nitrogen of well drained scrub soils prebum
and through 24 months postburn of scrub transects that burned in December
1986. Shown are means and 95% confidence intervals for the 0-15 cm layer (A)
and the 15-30 cm layer (B).
134
2.0 -
1.6-
1.2-
ZI 0.8 -N")
0Z
0.4 -
0.0
2.0
1.6
O)
o) 1.2E
V
Z 0.8I
0Z
0.4.-
0.0
//
, //Prebum
//
, //Prebum
NITRATEA. 0-15 cm
I0
I 1
6 12
AGE (months)
I
18I
24
B. 15-30 cm
I0
I6 12
AGE (months)135
I18
!2_r
Figure 36. Exchangeable ammonium-nitrogen of well drained scrub soils
preburn and through 24 months postburn of scrub transects that burned in
December 1986. Shown are means and 95% confidence intervals for the 0-15
cm layer (A) and the 15-30 cm layer (B).
136
45.0
40.0
35.0
30.0
E 25.0
20.0ZI
15.0-r"Z
10.0
5.0
0.0
25.0
20.0
15.0E
Z 10.0I
"i-X
5.0
0.0
//
, //Prebum
//
I //Preburn
AMMONIUMA. 0-15 cm
I I
0 6 12
AGE (months)
1
18
B. 15-30 cm
I
0 6 12
AGE (months)137
I
18
I
24
I
24
Figure 37. Exchangeable aluminum of well drained scrub soils preburn and
through 24 months postburn of scrub transects that burned in December 1986.
Shown are means and 95% confidence intervals for the 0-15 cm layer (A) and
the 15-30 cm layer (B).
138
50.0
40.0
.x 30.0
E
20.0
<C
10.0
0.0
11.0
10.0
9.0
8.0
"_ 7.0
#.) 6.0
v 5.0
4.0<I:
3.0
2.0
1.0
0.0
//
g
{3
' //Preburn
//
D
- o
- 1
, //Preburn
ALUMINUMA. 0-15 cm
I
0
I I
6 12
AGE (months)
1 I
18 24
B. 15-30 cm
01 I
6 12
AGE (months)
!3S
I I
18 24
Figure 38. Available copper of well drained scrub soils preburn and through 24
months postburn of scrub transects that burned in December 1986. Shown are
means and 95% confidence intervals for the 0-15 cm layer (A) and the 15-30 cm
layer (B).
140
0.4<)
0.35
0.30
0.25
E 0.20
0.15:3
¢,.)
0.10
0.05
0.00
0.30
0.25
O_ 0.20
O}E 0.15
0.05
0.00
//
COPPERA. 0-15 cm
I // 1 I 1 I I
Preburn 0 6 12 18 24.
//
, //Preburn
AGE (months)
B. 15-30 cm
I I I I I
0 6 12 18 24.
AGE (months)141
Figure 39. Available iron of well drained scrub soils preburn and through 24
months postburn of scrub transects that burned in December 1986. Shown are
means and 95% confidence intervals for the 0-15 cm layer (A) and the 15-30 cm
layer (B).
142
4.0.0
35.0
30.0
O)25.0
E 20.0
15.0
LI,,,
10.0
5.0
0.0
20.0
16.0
O)
12.0
v
8.0q)
i,
4.0
0.0
rl
m
//
i //Preburn
//
Ii
J.
I //Preburn
IRONA. 0-15 cm
I 1 I I I
0 6 12 18 24
AGE (monfhs)
B. 15-30 cm
I
0I I I I
6 12
AGE (months)143
18 24
Figure 40. Available manganese of well drained scrub soils preburn and
through 24 months postburn of scrub transects that burned in December 1986.
Shown are means and 95% confidence intervals for the 0-15 cm layer (A) and
the 15-30 cm layer (B).
144
3.0 ,
MANGANESEA. 0-15 cm
2.5
lm 2.0.X
0.5
0.0 , //Prebum 0
! I
6 12
AGE (months)
I18
I2'$
0.6 //
B. 15-30 cm
0.5
E 0.3V
0.1
0.0
T
, //Preburn
I
0I6
AGE
I ,, I12 18
(months)
124
145
Figure 41. Available zinc of well drained scrub soils preburn and through 24
months postburn of scrub transects that burned in December 1986. Shown are
means and 95% confidence intervals for the 0-15 cm layer (A) and the 15-30 cm
layer (B).
146
1.0
0.9
0.8
0.7
O.6
_0.5V
0.4.o-
N 0.3
0.2
0.1
0.0
0.60
0.55
0.50
0.45
0.40
0.35O)_: 0.30
0.25
c 0.20N
0.15
0.10
0.05
0.00
H
c]
m
w
i //Prebum
//
w
IIPreburn
ZINC0-15 cm
0I 1
6 12
AGE (months)
I18
B. 15-30 cm
I I I ! I
AGE (months)147
0 6 12 18 24
Figure 42. Precipitation from December 1986 to December 1988 during the
course of the scrub soil study. Data are monthly precipitation values from a
National Atmospheric Deposition Program rain station ca. 9.3 km south of the
study area.
148
ooo0Ob
JD
E(D
C 00 (D
Om
01
...O
EtO(D
d3
k\\\\\\\\\\\\\\\\\\\\ \
k\\\\\\
_\\\\\\\\\\\\\
k\\\\\\\\\\\\\\\\\\\\\\
k\\\\\\\\\\\\\\\\\\\\\\\"
N\\\\\\\\ \i
k\\\\\\\\\\\\\\\\\\\\\"
_\\\\\
• k\\\\\\\___
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N/LRA Report Documentation Page
1. Report No.
rM 103817
2. Government Accession No.
4. Thle and Subtitle
Dynamics of Vegetation and Soils of Oak/SawPalmetto Scrub After Fire: Observations FromPermanent Transects
7. Author(s)
Paul A. Schmalzer, C. Ross Hinkle
9. Performing Organization Name and Address
The Bionetics CorporationMail Code BIO-2
Kennedy Space Center, Florida 32899
12. Sponsoring Agency Name and Address
NASA/John F. Kennedy Space Center, FL 32899
3. Recipient's Catalog No.
5, Report Date
January 25, 1991
6. Performing Organization Code
BIO-2
8, Performing Organization Report No.
10. Work Unit No.
11. Contract or Grant No.
NAS10-11624
13. Type of Report and Period Covered
14. Sponsoring Agency Coda
MD
15. Supplementary Notes
_. Abstract
Ten permanent 15 m transects previously established in two oak/saw palmetto scrub standsqburned in December 1986, while two transects remained unburned. We sampled vegetation inthe > 0.5 m and < 0.5 m layers on these transects at 6, 12, 18, 24, and 36 months postbumand determined structural features of the vegetation (height, % bare ground, total cover). Weanalyzed vegetation data from each sampling by height layer using detrended correspondenceanalysis ordination. Vegetation data for the > 0.5 m layer for the entire time sequence werecombined and analyzed using detrended correspondence analysis ordination. Soils weresampled at 6, 12, 18, and 24 months postbum and analyzed for pH, conductivity, organicmatter, exchangeable cations (Ca, Mg, K, Na), NO3 -N, NH4-N, AI, available metals (Cu, Fe,Mn, Zn), and PO4-P.
Shrub species recovered at different rates postfire with saw palmetto reestablishing cover>0.5 m within one year, but the scrub oaks had not returned to preburn cover >0.5 m in 3 yearsafter fire. These differences in growth rates resulted in dominance shifts after fire with saw
17. Key Words (Suggested by Author(s})
Fire, Flodda, ScrubVegetation, Soil
18. Distribution Statement
19. Security Classif. (of this report) 20. Security Classif. (of this page)
Unclassified Unclassified
NASA FORM 1626 OCT 86
Unlimited, NTIS
-_'No. of pagesI49
22. Price